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Evolutionary roots of freedom To give up the illusion that sees in it an immaterial "sub stance" is not to deny the existence of the soul, but on the contrary to begin to recognize the complexity, the richness, the unfathomable profundity of the genetic and cultural her itage and of the personal experience, conscious or otherwise, which together constitute this being of ours: the unique and irrefutable witness to itself. Jacques Monod It is virtually impossible to discuss the cerebral foundation of liberty without dealing with the evolution of the brain. The reason is simple: The capacity of mammalian organisms to modify their environment by choice and to adapt to it by chosen means has grown enormously with the evolutionary growth of certain parts of their brain, the cerebral cortex in particular. Most relevant to our present discourse is the cortex of the frontal lobes. It is indeed a remarkable fact with a touch of cosmic irony that the science of evolutionary neurobiology, which can only "postdict" but not pre dict, has unveiled in the prefrontal cortex of man the seed of his future, the capacity to predict and to turn prediction into action that will impact on that future and on that of human society. The prefrontal cortex is the vanguard of evolution in the nervous system. Yet it is one of the latest cerebral structures to develop, in evolution as in the individual brain (Preuss et al., 2004; Rilling, 2006; Schoenemann et al, 2005; Sowell et al, 2003). Language and prediction, the two most distinctively human 28 Evolutionary roots of freedom functions that the prefrontal cortex supports, are anchored in the history of the species, as is the structure of the prefrontal cortex itself. In the human brain, the latter is tied to its evolutionary past and to the future it anticipates. Thus, while the human brain cannot predict evolution, it can predict the consequences of its actions, with them to predict and shape further actions in a continuous cycle, the perception/action (PA) cycle, which func tionally links the organism to its environment. The prefrontal cortex is the highest structure in that cycle, which integrates the past with the future - however near or distant either is - in the course of behavior, language, and reasoning. The PA cycle also has deep roots in evolution. In lower animals, earlier precursors of it mediate the adjustment of the organism to the surrounding world (Uexkull, 1926). The human brain, which sustains the PA cycle with the cortex, is the most complex adaptive system in the universe. It is an open system like all living systems (von Bertalanffy, 1950). As such, it is perma nently in quasi-equilibrium, but also in constant exchange with its environment to maintain that equilibrium. Thanks to its pre frontal cortex inserted in the PA cycle, the human brain, unlike any other, develops a prospective temporal dimension. Thereby, it makes advanced long-term adaptive changes in its environ ment. Furthermore, language endows the human brain with the ability to record those changes, to codify them, and to institution alize them. In short, the prefrontal cortex confers on the human brain the capability to predict and, accordingly, to preadapt. Lower brains have a measure of that capability, as well as certain primitive forms of communication with conspecifics that may be the ata vistic precursors of language. But the transition from the simian to the human in power of prediction and preadaptation, as well as communication, is so dramatic as to constitute a veritable quan tum leap. All relevant variables (complexity, time, "vocabulary," and so on) increase by several orders of magnitude. Along with it, variability increases immensely, so do the options for choice among alternatives. In fact, those comparative increases over other species are so large that the argument about functional homologies between humans and animals, even the great apes, 29 30 Evolutionary roots of freedom becomes well-nigh irrelevant. So does the discussion as to whether in evolution we are dealing with qualitative or merely quantitative differences in continua (Bolhuis and Wynne, 2009). With the advent of the human prefrontal cortex all the animal precursors ofcognition - intelligence and communication among them - open widely to a future agenda. This does not mean that the structure and workings of the animal brain are irrelevant to our understanding of the neuro biology of freedom. Quite the contrary; it is only in the brain of the animal, especially the nonhuman primate, that we can practi cally study the basic organization of knowledge, feelings, and values that give the human his or her freedom to make choices.1 In the animal brain we can investigate the mechanisms of the PA cycle behind choice, planning, decision-making, and the tempo ral organization of behavior. All of them are functions in which, as we will see, the prefrontal cortex plays a critical role. Those mechanisms constitute the underpinnings of human liberty, cre ativity, and their myriad expressions. Freedom, the capacity to choose between alternatives, emerges from the activity of cortical-cell networks of perceptual and executive memory, at the confluence of multiple converging inputs from past memory with multiple diverging outputs to future action. Freedom is a phenomenon of the brain's selection between those inputs and between those outputs for adaptive purposes. As a result of evolution and development, the cerebral cortex and freedom adopt in the human pivotal positions between an experiential convergent past and a divergent future of possibilities - and probabilities. 1 To be sure, no animal is amenable to the study of the semantic aspects of language and, least of all, to the neural mechanisms at their foundation. But many animal species lend themselves well to the exploration of the neural mechanisms of the temporal organization of information that language shares with all other cognitive functions. Those mechanisms are not directly accessible in the human brain, even with modern imaging methods. Needless to say. only in the absence of stress or pain are cognitive functions testable in animals; this imposes strict scientific - in addition to ethical - constraints on animal experimentation. Evolution of the cerebral cortex EVOLUTION OF THE CEREBRAL CORTEX Two approximate dates within wide uncertain ranges are especially relevant in the history of brain evolution: one 250 million years ago, and the other 250 thousand years ago. The first, in early Mesozoic, marks the appearance of the first mam mals, the second the appearance ofthe last hominid, Homo sapiens. The brains of fishes, amphibians, and reptiles were - and are covered by an evolutionally ancient cortex-like structure named the pallium (mantle). The pallium is divided into two compo nents, the hippocampal cortex situated next to the brain's mid line, and the piriform cortex, lateral to the hippocampal cortex (Figure 2.1). A third "pallium" will emerge between the two that in Homo sapiens will constitute 80 percent of the entire mass of the brain: the neocortex. B Dorsal cortex" Hippocampal cortex Piriform cortex Amphibians Reptiles Mammals Opossum Human Figure 2.1 Evolutionary development of the cerebral cortex. A: Lengthwise sections of the brains of four classes of vertebrates. P, pallium, generic name for cortex, both old and new (phylogenetically). From Creutzfeldt (1993). B: Crosswise sections of the brains of a primitive amphibian (Neclw-us), the box tortoise (Cistudo), the opossum (Didelphis), and the human being. From Herrick (1956), modified. 31 32 Evolutionary roots of freedom The neocortex or neopallium, the "new" cerebral cortex of the mammalian brain, remains physically wedged between the two ancient cortices, the hippocampus and the piriform cortex. The developing neocortex crowds the hippocampus toward the midline and the piriform lobe (piriform cortex and amygdala) toward the lateral underside of the brain. In later mammals, the neocortex grows heffily, pushing the two ancient cortices toward each other in the middle of the cerebral hemisphere. These two ancient cortices, even in the mammalian brain, preserve some of the functions they perform in primitive species: the sensing of life-sustaining signals, such as taste, olfaction, and spatial orientation. Additionally in primates, the hippocampus is involved in the acquisition and retrieval of memory, while the amygdala is involved in emotion, endowing memories with feeling. In the course of evolution, the neocortex, among all brain structures, increases the most in size, especially in primates. That evolutionary growth of the neocortex is exponential, out of pro portion with the growth of other structures. Further, the volu metric expansion ofthe neocortex occurs concomitantly with the differentiation of its cellular architecture, culminating in the human brain with a relatively large size and marked lamination of its cellular structure (six layers of neurons of different sizes, shapes, and densities). These changes, embedded in the human genome, appear to be the result of selective upregulation of gene expression2 relative to nonhuman primates (Preuss et al, 2004). Through those changes, and concomitant genetic mutations, humans have evolved mechanisms that allow them to overcome the physical constraints that impede the course of their own evolution (Krubitzer, 2009). Among the changes are those that take place in cortical architecture. Clearly, the evolutionary growth and differentiation of the neocortex has much to do with the increased ability to adapt to the environment and with 2 Upregulation refers to the increase in the capability of a gene to express cell products (e.g., specific proteins) in response to internal or external stimula tion, as, for example, an immune antibody to a new virus or an antitoxin to a new chemical agent. Evolution of the cerebral cortex the prolongation of life. In the human brain, that cortex has developed a large number of specialized areas to respond to all manner of sensory signals as well as to execute all manner of skillful movements. In brain evolution, the greatest neocortical expansion takes place in areas called "of association," which serve the higher cognitive functions; that is, those functions that deal with knowl edge and memory. Naturally, they deal as well with the neural transactions between the organism and the environment that depend on those functions. In the human brain, there are two separate cortical regions with areas of association. One is in the posterior part of the brain, extending over large portions of the parietal, temporal, and occipital lobes (PTO region), which con tains networks of knowledge and memory (cognits) acquired through the sensoiy systems. Those networks or cognits serve the highest aspects of cognition, including perception, language, and intelligence. The other associative region is the prefrontal cortex, the association cortex of the frontal lobe, which serves the executive aspects of cognition, especially the temporal organiza tion of actions in the domains of behavior, language, and reason ing. This "executive" cortex develops maximally in the human brain,3 where it occupies nearly one-third of the totality of the neocortex (Figure 2.2). Especially relevant to the development of the distinctive cognitive prerogatives of the human in language, planning, and Primarily on the basis of morphological imaging data (Semendeferi et ah, 1997), it has been argued that the prefrontal cortex does not evolve more, in proportion, than other cortical areas. Whereas this may be true volumetrically for the entirety of the frontal cortex, it does not take into account the fine structure of cells and fiber connections that characterize the prefrontal cortex perse. In the primate, anyhow, the frontal region that association nuclei of the thalamus innervate, which we call the prefrontal cortex, considerably exceeds in size the region they innervate in the posterior - perceptual - cortex (Jones and Leavitt, 1974; Walker, 1940). Further, on cytoarchitectonic grounds, the prefrontal cortex, which roughly corresponds to what Brodmann (1909,1912) called the regio frontalis (that is, all frontal areas minus areas 4 and 6), constitutes, by his calculations, 8.5% in the lemur, 11.5% in the gibbon and the macaque, 17% in the chimpanzee, and 29% in the human, of total cortex. On that basis alone, it seems legitimate to speak, figuratively, of an evolu tionary "prefrontal explosion" in the human. 33 34 Evolutionary roots of freedom Figure 2.2 Relative size of the prefrontal cortex with respect to total cortex in six animal species (marked by shading of external and internal hemispheric surface). PTO, parieto-temporo-occipital association region (posteriorassociation cortex). the exercise of freedom is the evolution of connections between prefrontal neurons and those of other cortical areas. Those connections, together with the neurons they link, constitute the essential components of the neural infrastructure of cognitive networks, and thus of all the cognitive functions of the cerebral cortex. Individual development of cerebral cortex The most rapid and efficient connectivity in the brain is that consisting of fibers surrounded by myelin,4 which constitute the bulk of the subcortical white matter and of the corpus callosum, the large commissure that connects the cortices ofthe two hemispheres together. It is the white matter, more than the gray - cellular matter of the cortex that increases the most in evolution. That is a clear indication of the vast expansion of the connective potential to form neural networks, which the human cortex requires in order to deal with the complexities of the world around it. Among all the connective tracts with which the human brain is endowed, probably none is more important for cognition than the one that links, in each hemisphere, the posterior (PTO), perceptual, cortex with the prefrontal, executive, cortex: the superior longitudinal fasciculus. The connections within that tract are bidirectional; in other words, they run in both direc tions, some PTO to front and others front to PTO. They constitute the backbone of the PA cycle, connecting posterior and frontal cortices reciprocally in tandem function. They will become essen tial for all kinds of temporally structured behaviors, where per ception will guide action, and vice versa, through the environment. Most importantly, they will be essential for the spoken language in dialogue, where the environment includes the interlocutor (Chapter 6). INDIVIDUAL DEVELOPMENT OF THE CEREBRAL CORTEX An old idea, first promulgated in 1899 by Ernst Haeckel (1992), is that ontogeny, the early development of an individual, recapitulates phylogeny, the development of the species. Indeed, many features of the individual nervous system of the human appear to develop in the same sequence they followed in the course ofevolution as a result of natural selection - that is, during the development of earlier animal species. Such features include, 4 Myelin is a white substance made of proteins and lipids that forms a sheath around the rapid-conduction fibers (axons) of the brain. It is indispensable for proper cortical integration and coordination. 35 36 Evolutionary roots of freedom for example, the prefrontal cortex, which develops late in evolution and does not develop fully until the third or fourth decade in the life of the individual. Haeckel's recapitulation idea is basically flawed in one important respect. Whereas natural selection follows a passive process based on random genetic variance and mutation, ontogeny follows an order established by biological clocks that determine gene expression and molecular-enzymatic change jointly with environmental influences from point of conception onward. Nevertheless, Gould (1992) attempts to reconcile the two trends, phylogenetic and ontogenetic, by proposing the concept of "heterochrony." Heterochrony would simply signify the change, in the development of the individual organism, of the relative rate and timing of appearance of char acters already present in ancestors. In other words, ontogeny by its own clock would compress, extend, and retime the results of evolution. The concept of heterochrony would legitimize the predic tion of ontogeny based in part on the postdictable evolution of traits, but that would still leave that prediction subject to the uncertainties surrounding lineage and ancestry in evolution. In any event, it would be wrong to deny or obscure the evidence that the development of the human brain to adulthood entails the development of social traits already present in the behavior of most animal populations. Among those traits are the inborn tendencies to affiliation, trust, group protection, and hierarchical social structure. Consequently, it is fascinating to study brain development in the attempt to glean how it relates not only to the development of cognitive functions that are distinctly human, such as prediction and language, but also to the develop ment ofsocial dynamics already present in ancestral populations. Both issues bear on the roots of freedom. There is an added incentive to study the ontogeny of the brain from an evolutionary point of view. It is now increasingly evident that, just as in the evolution of species and traits, in ontogeny the development of the brain's features and functions is the result of dynamic interactions between the elements of biological populations: gene populations, neuron populations, Individual development of cerebral cortex synaptic populations, network populations, and nerve fiber populations. The neonate comes into the world with the structure of the cerebral cortex practically complete, with all its principal ele ments in place. The neocortex, the "new" cortex in evolutionary terms, is already characterized by its laminar structure, the pres ence in it of the major types of nerve cells, synapses, and other contacts between cells, as well as the major excitatory and inhib itory chemical neurotransmitters.5 In quantitative terms, how ever, certain fluctuations occur over time in the relative amounts of those elements. There are periods of exuberant pro duction of neurons followed by periods of attrition in their num bers. The same may be said for synapses and other elements of cellular architecture. From the beginning, however, there is a gradual and more or less continuous increase offiber connections between cells in most all layers of the neocortex (Figure 2.3). This increase in connectivity persists into adulthood and is most mani fest jn the myelination - covering with myelin - of long fiber connections between cortical areas. This translates itself into general increments of cortical white matter even in the presence of some relative decrements in gray matter. The age-related increase in cortical connectivity is critical for cognitive development, and, therefore, for the development of free will. Connectivity is essential to a relational code, such as the code of cognition and of the cognits of memory and knowl edge.6 Cognits are defined by relationships between elements (neurons or assemblies of neurons) that represent discrete com ponents of a memory or item of knowledge (Chapter 3). Byvirtue of the combinatorial power of connections, an also discrete num ber of neuronal assemblies can encode, by combination and per mutation, an almost infinite number of different items of The most important excitatory neurotransmitters are glutamate, norepi nephrine, serotonin, dopamine, and acetylcholine (Siegel,1999).The concen tration of each of these varies somewhat from area to area. By far the most important inhibitory transmitter is the ubiquitous gamma-aminobutyric acid (GABA). A study of brain connectivity by reliable neuroimaging reveals the age-related enhancement of cortico-cortical connectivity in children performing cogni tive tasks, such as listening to stories (Karunanayaka et al., 2007). 37 38 Evolutionary roots of freedom 3 mo. 6 mo. 15 mo. 24 mo. Figure 2.3 Development of neurons in the human cortex. Top: Prenatal period, from 10.5 weeks to birth. From Mrzljak et al. (1990), with permission. Bottom: Postnatal, at 3, 6,15, and 24 months. From Conel (1963), with permission. memory or knowledge. The same can be said for the cognits and the relationships between them, which can be part of larger cognits. In fact, it is that combinatorial power of connections that gives us the individuality of our memory and of our actions. The existence of more synaptic connections than neurons can provide the executive cortex, especially the late developing cortex, with enormously diverse inputs and outputs. Thus the prefrontal cortex can thereby give rise to - i.e., organize - an immense number of alternative actions. Given that cortical con nectivity increases with age at a higher rate than brain mass, it is reasonable to conclude that the options ("choices") of both inputs Individual development of cerebral cortex to and outputs from the executive cortex also increase with age at a very high rate. The same is true for freedom, which is in essence the capacity of that cortex to selectively favor or bias inputs and outputs. The best direct evidence of the age-related increase and reinforcement of cortical connectivity is the age-dependent mye lination of long cortical nerve fibers. We have known for over a century, since the seminal work of Flechsig (1901), that around the time of birth cortical myelination follows a certain chrono logical order, which can be discerned by histological fiberstaining methods (Figure 2.4). The first to myelinate are the sensory and motor areas of the cortex. From then on, myelination takes place in progressively higher areas ofassociation.7 The last areas to fully myelinate are those of the posterior and frontal association cortices (white in Figure 2.4). Thanks to modern scan ning methods, we now know that the prefrontal cortex does not reach full myelination until the third or fourth decade of life (Sowell et al, 2003). The implications of this fact are profound, especially as they relate to cognitive maturity and, of course, freedom of action and responsibility for it. Assuming that the degree of myelination is related to neural maturation generally, and assuming further that neural matura tion is related to psychological maturation, it is reasonable to speculate on the age-related neural constraints of psychosocial development. For example, it seems more than just possible that much of the turmoil of adolescence is caused by an imbalance between the two sides, emotional and cognitive, of the PA cycle. On the one side is the input from emotional centers under the onslaught of massive hormonal changes; add to that the exigen cies ofgratification in the presence of still immature principles of behavior. On the other side is an immature prefrontal cortex 7 The staining of myelin in anatomical specimens of the cerebral cortex is not a simple matter. It is a laborious technique subject to errors, some of which have been pointed out by seasoned neuroanatomists as possibly having distorted Flechsig's original observations. Nonetheless, whereas some quibble about the precise order of myelin formation as portrayed in Figure. 2.4, there is general consensus on the conclusion that the process commences in primary sensory and motor areas and continues through the association cortex. 39 40 Evolutionary roots of freedom Figure 2.4 Numerical order of myelination of areas of the human cortex, according to Flechsig. Primary sensoiy and motor areas (low numbers, in black) myelinate first; association areas (high numbers Inwhite) myelinate last. From Bonin (1950), modified. ready for physical action without the capacity, yet unavailable, for reasoning or good judgment. The result is self-oriented and self-adjudicated liberty with minimal responsibility, characteris tics of the typical teenager. Neural Darwinism By age 20, freedom has in most individuals acquired a social dimension, and with it the social responsibility that constrains, or rather complements individual freedom, is nearing its adult plateau. The third decade oflife calls for sharp cognitive decisions on one's future. By then, full maturity is reaching the highest areas of cortical association. With it the brain reaches the peak of inventive capacity and imagination. With the maturation of the prefrontal cortex in particular, language and the capacity to pre dict expand, and with them the capacity for social planning with common purpose. Those capacities will lead to decisions at higher level, together with more freedom to lead others to greater enterprises - educational, scientific, artistic, legislative, sporting, and so on. It is the time when careers get started, superior studies undertaken, and plans made for emotional, professional, or social associations with others.8 With further cortical maturation, more elaborate, complex, and abstract cognits are acquired and con solidated in the cortex as part of the individual's experience. These cognits include, among others, principles of altruism and social justice. NEURAL DARWINISM The brain is essentially the organ by which the animal, through sensing and acting, adapts to its environment. As such, the brain develops modes to adapt to that environment that are similar, if not identical, to those that guided evolution. A princi ple of development that applies to ontogeny as well as evolution is natural selection. To be sure, natural selection works for grow ing individuals in different ways than for growing species. But its s Nonetheless, it is somewhat simplistic to ascribe the acquisition of any given social trait to narrower chronological age spans. In the first place, any plfenotypical trait at any age is the result of the interaction of genetic factors with environmental factors. There is individual variance in both sets of factors. Environmental factors intervene at different ages depending on internal con ditions, such as hormonal levels. That interaction, in turn, leads to changes in social interaction. To this we have to add the imponderables of individual differences in nervous and hormonal maturation. The result is a long and complex series of PA cycles between the individual and society that defy precise chronological bracketing. 41 42 Evolutionary roots of freedom adaptive results are similar, and to some extent parallel and synergistic. Something radically new, however, takes place in the human brain that is unprecedented in prior evolution. Largely on account of the extraordinary evolutionary growth of its pre frontal cortex, the human brain "opens" to the future. Selection is no longer between items of information or action that have occurred in the past or are to occur in the immediate future. The cerebral cortex of the human has become predictive. With that change, selection can be made between anticipated options of percept and action to occur in the future. Caution is needed here, however. The anticipating agent is not consciousness, the "ego," or some other subjective entity. It is the cortex itself, which by predicting becomes preadaptive. Our will is as free as our cortex is free to select future actions and prepare for them. It is as if human development had forced a Copernican shift on evolution, from the past to the future - a shift, nonetheless, that does not change the basic principles of selection, variance, and probability that move evolution under the umbrella of adap tation. There are, however, two new principles that appear with the human's prospective adaptation: teleonomy and affordance, which I shall discuss later. Under the title Neural Darwinism (1987), Edelman proposed a new theory that applies evolutionary principles to the individual brain. His theory of neuronal group selection (TNGS) relates to the formation or modulation of brain circuits as a result of the sensoiy contacts of the organism with its environment. Originally, the cor tex and its link to the outside - that is, the thalamus - come into this world with a genetic endowment of what he calls a primary reper toire of interconnected neuronal groups in the two structures. Through interactions of the animal with the environment, and the neurobiological mechanism by which "cells that fire together wire together," a secondaiy repertoire of cell groups will be formed. This secondary repertoire will emerge at the expense ofthose cell groups not selected, which will wither away - in correlation with the observed postnatal attrition of cells and synaptic contacts. The selected groups will self-reinforce their connections by circuit re entry - output returning as input. In this manner perceptual Neural Darwinism experience will be acquired and registered in thalamic-cortical cir cuitry. A similar argument can be applied to the dispersed neuronal groups of the cerebral cortex, which, if they fire together, will wire together into cognitive networks. Re-entry is the universal consolidator and activator of those networks.9 Regardless of the precise role of genetics in phylogeny and ontogeny, the fact remains that in the nervous system certain principles apply to both. These principles generally apply to the adaptation of all biological organisms to their environment and are unmistakably present in both phylogeny and ontogeny. They include variance, selection, and probability. In the human organ ism, with its prospective properties, we have to add teleonomy and affordance (below). Variance is the essential precondition of evolution. It is in response to variance, whether in random gene mutation or in environmental change, that natural selection occurs. Traits, fea tures, competitive advantages, and so on are selected by nature (note the passive voice) to adapt the organism to its environ ment; the adaptation occurs at the level of the population of the species - indeed, evolution is a population phenomenon - with the end result of furthering survival and procreation. Much of the selective adaptation, of course, takes place in the nervous system. Variance in the nervous system, at the interface of the organ ism with the environment, also serves selection and adaptation in the individual. Here, however, the selection - say, between sensory inputs and between actions - is active - that is, from the organism outwards. As in evolution, the selection is adaptive, but now the organism exercises it actively on the world. Evolution has in fact given the individual the means to do it by itself. As in evolution, selection serves the adaptive ends of the population, beginning with brain cells and circuits and extending to the social order. 9 The Darwinian aspects of the TNGS can be criticized by the same argument that distinguished the role of evolution from that of ontogeny in the forma tion of neural structure. Even Gould's concept of heterochrony does not quite reconcile the two. Thus, in the defense of neural Darwinism, the latter has been simply referred to by some as a metaphor of the evolutionary process. Nobody disputes, however, the critical importance of re-entry in the structural and functional development of the cerebral cortex (Edelman, 1987). 43 44 Evolutionary roots of freedom Selection takes place in all the interactions ofthe brain with the milieu, both internal and external. It serves the related purposes of economizing resources and increasing efficiency. On both counts, selection works on all percepts and all actions. In those two domains, selection performs two separate but syner gisticadaptive functions: (1) categorizingand (2) discriminating. Perception is the categorizing of the world that surrounds us (Hamad, 2005; Hayek, 1952). We perceive, that is, categorize objects by virtue of their common features and the relations between their parts (next chapter). The identity of an object stays the same despite wide variations in size or other features of its parts, provided that the relations between some of those parts stay the same ("arose is a rose is a rose," despite differences in color, shape, size, or fragrance). This is the fundamental psy chological principle of perceptual constancy, which says that we perceive an object as the same regardless of variations in size, perspective, color, shape, and so on. In our daily life we continuously perceive - mostly uncon sciously - the objects and events around us by classifying those objects and events into categories, and by matching them to previous experience - that is, by matching them to established cognits in our cortex. Conversely, we distinguish and discriminate between objects and events as we concentrate on their individual features (a yellow rose is different from a red rose). Categorizing and discriminating are tandem functions in the establishment of sensoiyorder in our cortex(Hayek, 1952). Theyguide not only our ordinary life but also our scientific endeavors. Deduction and induction, generalization and analysis, depend on them. The two selective functions of categorizing and discriminat ing also operate on the side of action. Now the categorizing principle is the purpose or goal of the action. Many possible movements can lead to the same outcome. We may call this function "action constancy." At the same time, large goal-directed movement is composed of small subcomponents to attain differ ent subgoals on the way to a major goal (Bernstein, 1967). Clearly, the categorizing of either perception or action in the nervous system cannot be accomplished without something akin to the principle of degeneracy of Edelman (Edelman and Neural Darwinism Gaily, 2001). In essence, degeneracy refers to the fact that in the brain, as in other complex systems, multiple inputs can lead to the same output.10 No organism could survive without it. Degeneracy, or something like it, is at the root of perceptual and motor constancy. The cerebral cortex is permanently in a state of internal change, yet that change tends to equilibrium at some point in the future. The billions of neurons concomitantly active in the vigilant cortex, whose electrical activity is characterized by "desynchron- ized" rhythms,11 would bombard sensoiy and motor centers with such a diverse flow of impulses that those centers would easily be led to chaos. In the absence of degeneracy, in other words, in the absence of the capacity to generalize across inputs or outputs, no stable perception or action would be possible. In the state of attention, a cognitive function that is selective by definition, selection is evident in the two components of the PA cycle, perception and action. Attention could rightfully be considered the mother ofall cognitive functions. It selects certain percepts, memories, motives, and actions at the expense of all others, which are suppressed and inhibited (Fuster, 2003). But again, this happens with or without consciousness, though con sciousness is a constant phenomenon in the most demanding selec tions. It happens as a result of the internal dynamics ofthe cortex, without the need for a central executive. Certainly, the prefrontal cortex serves attention, but merely as the mediator of selective perception or action. Any control from this cortex over attention and over other cortical regions - derives exclusively from its dynamic involvement in the PA cycle (Chapter 4). In the human brain, selection - especially selective atten tion - reaches into the future. The cerebral cortex selects percep tual and executive cognits for goal-directed prospective action, while other less relevant cognits are inhibited. Here evolution Conversely, in sensoiy discrimination and in discriminating action, one input, depending on certain features of it. can lead to different outputs. "Desynchronization" is a characteristic of the cortical electroencephalogram (EEG) in the awake state. It probably reflects the re-entrant activation of multiple cognits. each at its own frequency range or "spectral fingerprint" (SiegeletflL 2012). 45 46 Evolutionary roots of freedom has endowed the human brain with the ultimate cortical appara tus to do that prospective selection, in the full sense of the word "ultimate": the prefrontal cortex. It is with the prefrontal cortex that the human brain acquires its freedom to set goals and purposes. In science, teleology is a dirty word. It is also a logical incon gruity, because it implies the temporal inversion ofcausality, which is just about the worst possible anomaly in scientific discourse. Yet it is extremely compelling to attribute to the prefrontal cortex a teleological function. Things appear to happen in that cortex because of a future event, whether that event is a course of future action, a goal, a reward, or the answer to a request from somebody. Is a future event the cause ofpresent action? Here the laws of physical causality would seem to be turned backwards and upside down. That is just an appearance, however. With the advent of the prefrontal cortex, goals and purposes have entered the agenda of the brain. On close examination, the teleological paradox dis solves before our eyes, because the cause of future action is firmly anchored in the past. That past, in the brain, consists of evolu tionary and individual memory in the form of established drives and imagined cognits of the future; it is the "memory of the future" (Chapter 5). A better word for that land of teleology is teleonomy (Monod, 1971), which, in essence, is a critical dimension of liberty, perhaps in effect its most decisive dimension. Teleonomy has been identified with life, as its future preservation is the first objective of life itself. Out of the "night of evolutionary time," Homo sapiens emerged, the end result of countless interactions of countless organisms with their environment. At the crux of those interactions was a long and silent cycle of mutual influences between genes and "environmental demand." By a process that we do not understand, and probably never will, the brain of Homo sapiens ("knowing man") acquired the means to foresee and foretell the "demands" of the Umwelt (the world around) and to change them in order to better adapt to it during his life and that of his descendants. The PA cycle grew in complexity. Surely there is a rudiment of it in other primates, but no more than a rudiment (next section). With the human brain, the Neural Darwinism temporal period of the cycle increased by several orders of mag nitude. So did the complexity of the agent, now the brain itself, and the complexity of the environmental information it was able to handle, and to predict. The prefrontal cortex devel oped as the supreme neural predictor at the top of the cycle. Language is an immense elaboration of animal communica tion. It also emerged from the expansion of the prefrontal cortex with its temporal organizing properties. Language became a marvelously suited means of closing the PA cycle between the brain and the environment in the service of the self and others. With language, the human brain became capable of formulating prob abilities of future causality, to bias favorably those probabilities with logic (Greek, logos, word) for the benefit of the self and others, and to record changes, both past and projected into the future. Language adds another decisive dimension to freedom (Chapter 6). The psychologist Gibson (1977) coined an interesting term, ajfordance, which fits perfectly in the human PA cycle, especially in its future perspective. Affordance is a quality of an object or environment that allows a subject to perform an action. Thus, affordances are action possibilities that the world offers to the individual. I think the term and its definition are useful here, but that definition should be expanded, as the human being is capa ble of creating affordances and of projecting them on the environ ment. New affordances can thus emerge from perception of the environment in the light of prior experience - that is, in the light of established cognits. The invented new cognits, projected on that environment, can thus be incorporated in the PA cycle and guide the action to the adaptive manipulation of its objects. Affordances, therefore, are another means by which the prefron tal cortex can imagine ("memorize") future action. Affordance is yet another decisive dimension of freedom. The human being is unique in that it can freely create his or her own affordances. As we will see in the corresponding chapter (Chapter 6), language is an extraordinarily fertile creator and vehicle of affordances. The prefrontal cortex, the most advanced product of evolution in the human brain, is the supreme enabler of both language and affordances. 47 48 Evolutionary roots of freedom THE TWO TEMPORAL FACES OF LIBERTY Just as Janus the Roman god had two faces, one looking backwards and the other forwards, liberty has two temporal perspectives, one lookingto the past and the other to the future. The analogy hasbeenaptly applied to the conscious experience of time, which has been called "mental time traveling," between the past and the future. Now, applying it to liberty requires some explanation. Of course, we are not free to change the past, for the past is "done." But the Janus analogy is valid with regard to freedom because of the simple fact that, while we are not free to change that past,we are free to choose parts of that past to make informed choices for the future. Further, a chosen action is not only based on prior experience, but it also engenders new experience to inform future choice, thus completing the PA cycle. In effect, however, the lion's share of prior experience at the base of our freedom to make choices is not exclusively our own, but belongsto the entire human species;it is made of our sensoiy and motor systems. That common "experience" is built in the genome and finds its expression in the physical anatomy of those systems. For that reason, I call the structure of those sys tems "phyletic memory." It is genetic memory in the form of the nervous structures and mechanisms that are essential for ecolog ical adaptation. Phyletic memory includes the peripheral recep tors of primary sensations and the generators of elementary movements for nourishment and defense.12 It should be apparent therefore that the term phyletic memory is more than a figureofspeech, in that the memory it carries in its very structure is the collective experience of the species in deal ingwith the physical environment. It can be legitimately called memory because it consists of "stored" information that, after 12 Tocallthe sensoryand motor systems"structural memory" makessense only in evolutionary and ontogenetic terms. Surely it is fundamentally genetic memory in the sense that the structure of those systems is encoded in our genes.Even before birth, however, it is subject to environmental influences upon the phenotype of those two systems. The two temporal faces of liberty critical neonatal periods of "rehearsal,"13 is retrieved and utilized with every act of perception or overt action. The primary sensoiy and motor cortices are part of that memory and comprise the indispensable interface between evolutionary and individual memory.14 It is through the functioning of phyletic memory in those cortices that individual memory, perceptual and executive, is formed and deposited in the cognitive networks (cognits) of the cortex of association. Liberty rests on the potential selectivity of those individual associative cortices, and it is logically constrained by sensoiy or motor handicaps that affect their phyletic base. In any event, it is the cortex as a whole that makes the choices that are the essence of individual liberty, not an extracortical or extracorporeal entity that we can identify as a choos ing, deliberating, and willing I. The I is nothing other than the cortex, selecting between inputs, some from the past and others from the present, in order to select outputs of adaptive action.15 Thus, individual selection is no longer the natural selection that served the population in evolutionary time; however, the cogni tive choices of the individual crucially depend on that phyletic history. The selective cognitive networks of the cerebral cortex are constantly under the influence from another store of phyletic memory situated deep in the interior of the cerebrum: the 13 Duringcertain criticalperiodsshortly after birth, the senseand motor organs have to be utilized to become fully functional from then on. Animals that through those periods have been deprived of visual or auditory stimulation become permanently impaired, visually or auditorily, because of faulty cort ical development (Hensch, 2004). M The primary motor and sensoiy cortices, while being the lowest and most basic levels of phyletic cortical memory, are not the lowest stages of inherited evolutionary sensors and effectors in the central nervous system. Arguably, we have to descend to the spinal cord and to the nuclei of the autonomic nervous system in the brainstem to find them. For it is in these structures where lie the centers of reflex activity that regulate the most primitive, innate, defensive, and nurturing mechanisms with which we adapt our organism to the internal and external milieus. 15 Each cortical choice is the result of a massive process of computation of inputs from many sources upon the association cortices, and ultimately the prefrontal cortex. An important point here is that the prefrontal cortex enables and mediates the action within the PA cycle, but is not the sole generator of that action. 50 Evolutionary roots of freedom emotional or limbic system or brain. The limbic system consists of an array of interconnected neural masses and nuclei of ances tral phylogenetic origin that critically intervenes in the imple mentation of instinctual drives and emotional responses of the organism to the environment, internal and external. The fore most components of the limbic system are the hypothalamus, the amygdala, and the hippocampus. The first two are implicated in all instinctual behavior (feeding, sex, flight, defense, and aggres sion), as well as in the acquisition, maintenance, and retrieval of emotional memory - that is, the memory of likes and dislikes, love and hate, reward and punishment, pleasure and pain (Denton et al, 1996; Hess, 1954; Pessoa and Adolphs, 2010). The hypothalamus and the amygdala are also directly or indirectly connected with the autonomic nervous system and hormonal systems, which play important roles in visceral control and emo tion (Buijs and van Eden, 2000). The hippocampus is a transitional ancestral structure situ ated between the limbic brain and the neocortex. It is anatomi callya portion of ancient cortex, folded onto itself and tucked in the middle of the cerebral hemisphere on each side. In lower mammals, it performs vital functions in olfaction, touch, and spatial navigation.16 In primates, especially the human, the hip pocampus plays a critical role in the acquisition, consolidation, and retrieval of memoly of any modality (Squire, 1992). Adjacent to the limbic brain are the basal ganglia (Aliens Kappers et al, 1960), a conglomerate of neural structures, also of early phylogenetic and ontogenetic development, that critically intervene in voluntary, reflex, and automatic motility. Connective loops that link the motor cortex, the cerebellum, the thalamus, and the basal ganglia mediate the timely execution 16 Ithas always beensomewhatofa mysterywhythe hippocampusof rodents is so critical for these three functions, whereas the human hippocampus is only marginally involved in them. The reason, in my opinion, is because the hippocampus of the rodent (which is the most advanced cortex the rodent has) is in charge of the three functions inasmuch as they are essential to the animal's survival, whereas in the human, those functions have migrated to higher cortex for more flexibility and range of adaptation to a more complex environment. The two temporal faces of liberty of voluntary - as well as automatic and well-rehearsed - motor sequences (Alexander et al, 1992; Kreitzer and Malenka, 2008). In conclusion, those internal parts of the brain, together with primary sensoiy and motor cortices, in the aggregate, con stitute the cerebral ground layer of phyletic memory. That struc tural memory, including predispositions to action in the basal ganglia, is the primordial neural apparatus for adaptation to the environment. That environment includes the internal milieu, which was selected in evolution to fulfill the most immediate exigencies of survival and procreation of the species. Freedom will rest on that fund of evolutionary experience, while remaining also constrained by it. For, no organism, includ ing the human, can surpass the limits imposed by the sensoiy and motor capabilities it inherits. In other words, that heritage after critical postnatal periods - limits our senses to light, sound, touch, olfaction, and taste within certain ranges of frequency, intensity, and chemical composition. It also limits the range of angle, direction, and bearing of each of our joints and limbs. Thus, considering the genetic endowment ofour motor systems, there are physical limits to the actions we can execute with those systems. This remains true even after full development and physical education. Freedom will not only rest on, but also emerge from, that primordial structural memory of sensoiy and motor systems, visceral control systems, and emotional systems. The most imme diate conditions permitting its emergence are the variance and plasticity of those systems, and, most critically, the cognitive networks of the cortex of association. In the individual human, those systems and networks will provide the essential inputs to the PA cycle, which moves us from one choice to the next, from one decision to the next, and from one objective to the next. The aggregate ofthose inputs constitutes the experience, phyletic and personal, on which liberty is based. It is the major share of the retrospective aspect of liberty. How free are our choices of retrospective memory? They are, and must by necessity be, only relativelyfree because they are not free of physical and psychological constraints. Regardless of those constraints, our freedom depends on, and is directly 51 52 Evolutionary roots of freedom related to, the availability of alternatives, whether we are aware of them or not. We experience freedom even though and in part because - we are unaware of the degree to which those alternatives steer our actions. This is not a blanket endorse ment of determinism. On the contrary, having decoupled cortical choice from consciousness, freedom asserts its independ ence. Determinism is tempered in the variability and randomness of a complex adaptive system such as the human cerebral cor tex.17 But here the concept of "cortical choice" needs to be qualified. Choice implies alternative. But the alternatives of informa tion that reach the cortex to literally inform action do so with different degrees of intensity (synaptic strength) depending on a variety of internal and external states or circumstances. Some alternative inputs will be in conflict with one another; others will potentiate each other. The inputs that prevail in the decision to act will be the probabilistic result of competition or summa tion of synaptic "weights," which will sway the cortex to one action or another. Probability here is to be understood in the Bayesian sense, in this manner applicable to the state of the evidence or knowledge contained in any given set of inputs.1S Ultimately, the weight of each input on a decision will be "deter mined" according to an estimate of probability based on prior knowledge and bearing on the synaptic weight of that input.19 As we have seen, alternatives of input will arrive in the prefrontal cortex from many sources, some cortical and others subcortical. Alternative sources of input will multiply if the The concept of open adaptive system was first proposed by von Bertalanffy (1950). the father of "general system theory" (GST), to account for the dynam ics of organisms - biological and sociological - that tend to equilibrium or steady state in the face of perturbations.They do it by use of self-correcting feedback, among other mechanisms. Clearly, that cyberneticconceptapplies to all self-regulationin the nervous system, and of course the PA cycle. Bayesian probability, as distinguished from "frequential" probability, is the logicbased on uncertain statements about a hypothesis and liable to change by the acquisition of further data (jaynes et al., 2003). The expression "synaptic weight" is meant here to encompass not only the strength of present or potential electrochemical transactions at the mem branes of nerve cells, but also the numbers of axon connections and other input fibers arriving at those cell membranes. The two temporal faces of liberty actions informed by those inputs are complex and high in the hierarchy ofthe PAcycle. Bycontrast, inputs of subcortical origin, at the foundations of phyletic memory, will be simple and straightforward, such as the urge to eat, fight, or mate. They will arrive to the cortex by fiber paths funneled through the orbital prefrontal cortex, which collects information about rewards with biological "valence." Rarely, except in a sociopath, will those impulses come to the cortex unaccompanied by inputs from cortical networks representing social, ethical, and esthetic prin ciples. Hence the formalities of dining, sport competition, and courting. Those natural impulses will come to our cortex also accom panied by inputs from the cognitive networks that store our episodic and semantic memory. Those too will be subject to internal "competitive bidding" in the higher levels of the PA cycle. Consider, for example, the many inputs that, in addition to hunger, inform our choices of restaurant and menu: personal experience, counsel from friends, cost, means of transportation, parking availability, ethnic-food preference, and so on. All will weigh on the choices. In sum, on the retrospective side of liberty are the inputs to the orbital prefrontal cortex from the internal milieu and its limbic sources. Concomitantly, inputs from the cortex at large converge on the cortical lateral convexity of the frontal lobe, conveying to it information from the cognitive networks of knowledge, personal memory, and social values. The prefrontal cortex will reconcile and prioritize those inputs before each deci sion. Completing the PA cycle, the prefrontal cortex will also reconcile and prioritize the consequences of each action for fur ther action. The reconciling and prioritizing will be done in and by the cortex itself. Those operations will take place in a multi variate and probabilistic environment of synaptic connections of widely differing and variable weights. It is the relative internal configuration of those weights with respect to one another that will lead to one alternative of action or to another. The root of the decision, therefore, is to be found in all its antecedents and the relative synaptic weights of each of their respective neural foundations. 53 54 Evolutionary roots of freedom EVOLUTION OPENS MAN AND WOMAN TO THEIR FUTURE Inseparable from the retrospective aspect of freedom is its prospective aspect, which willbe further discussed in Chapter 5. The two are two sides of the same coin, namely, of the dynamics of the PA cycle, which by definition has a past and a future alternating with each other. Both are anchored in phyletic memory, from which the selectivity of individual human cognition flows. Arguably, action precedes perception, in phylo geny as well as ontogeny. Just as evolution selects from genetic variance "actions" that are adaptive for the population, the human infant enters the world palpating it to select certain "adaptive" stimuli within it. It is by haptics - active touch - that the newborn finds the mother's breast. Crying, which is another phyletic action, will rapidly join haptics in the PA cycle of infant nourishment.20 With the evolution ofthe cerebral cortex in general, and the prefrontal cortex in particular, the PA cycle of the human begins early in life to workat higherlevels of complexity for the benefit of the individual and, eventually, society as well. Liberty will emerge from the expansion of both the fund of alternatives of information available to the cortex and the alternatives of action to which that information can lead. Among the alternatives of action, none is as relevant to liberty as those that evolve in the fields of cognition most charac teristic of the human being: planning and language. Here we have to address a question that often arouses endless controversies. Are there precursors of planning and language present in the animal kingdom before man? The most immediate - and least controversial - species to queiy in this respect are the great apes. Away to assess the ability to plan behavioris to examine the ability to use tools, since tool use invariably implies a certain sequence of personal actions that is reasoned, goal-directed, and 20 Later, "babbling" and the rudiments of languagewill be incorporated in the PA cycle betweenthe childand the environment presidedover bythe mother (Chapter 6). Evolution opens us to our future not innate. For several years around the time ofWorld War I, Kohler (1925), a noted gestalt psychologist, studied meticulously the reason ing potential of chimpanzees in a colony of such animals he main tained on one of the Canary Islands. One of his smartest subjects. Sultan, learned to stack boxes on winch to stand and to use poles to reach high-hanging bananas. From this Kohler concluded that the animal was able to reason to some degree and to use intermediary objects to attain spatially distant goals. Whether Sultan planned actions or used tools in the strict sense of these words has been extensively debated. It is unques tionable, however, that the animal was capable of a degree of prospective and purposive reasoning. It is also unquestionable that he learned or had the intuition to use "tools" to attain his goal. Lower primates are capable of doing both, though to a lesser degree. Later studies, furthermore, have shown that the great apes are capable of cognitive "time-traveling" and foresight, as they can display sequential actions to reach goals that are distant in space and time (Osvath and Osvath, 2008). The argument that all those operations are the result of associative learning is idle, because there is no foresight or tool use of any kind without some degree of prior associative experience. In fact there is no liberty to act reasonably one way or another without prior empir ical knowledge of context and consequences. Thus, there is rudimentary planning and foresight in the earlier primates. However, because humans exceed those capa bilities by many orders of magnitude, it has been mistakenly argued that those capabilities are the exclusive patrimony of our species. Similar arguments and counterarguments have been made with respect to language, albeit usually with more vehemence on both sides. The most obvious question in this respect is whether the vocalizations of nonhuman animals qual ify as evolutionary precursors of language. The answer is yes, insofar as those vocalizations consist in means of communication between conspecifics, like language. Another, more relevant, question is whether animals possess some primitive form of language. The answer here is definitely no, as animals, while able to communicate with symbols, cannot communicate with logical reasoning, which is an essential attribute of language. 55 56 Evolutionary roots of freedom With language and the enormous expansion of the cerebral cortex that goes with it, comes the unique tool that more than any other allows humans to fashion their future: speech. This is the supreme maker of affordances, a la Gibson (1977). It is not a coincidence of nature that that "ultimate tool" (Greenfield, 1991) develops together with a region of the frontal cortex that is heavily involved in tool malting and utilization.21 Nor is it coincidental that the entirety of the frontal cortex, where that area is located, serves not only speech but also, more broadly, what Lashley (1951) called the "syntax of action." Indeed, by speech, evolution comes around to furthering an unwritten "pur pose" of the evolution of the species: the freedom to ensure its own survival. For it is by the written word that the human race codifies the liberty of its progeny. CONCLUSIONS Our freedom and ability to shape our future are the ultimate offspring ofthe extraordinary evolution of the human brain. Both freedom and creativity have their most recent evolutionary root in the prefrontal cortex, the latest domain of the cerebral mantle to attain structural maturity, in evolution as in individual development. Critical for both human prerogatives, freedom and future, is the rich connectivity that develops between prefrontal cell pop ulations, as well as between them and those in other cortical regions. Because of the inherent synaptic plasticity of that con nectivity, cognitive networks (cognits) will be formed by life experience in the associative cortex, which will codify the mem ory of the individual and inform his decisions to reach his goals (Chapter 3). Freedom flourishes with the capacity of the cortex to choose between memory networks and between action networks in the pursuit of chosen goals. In all instances, the pursuit of a goal takes 21 The area in question includes Broca's area and a large portion of the premotor cortex, both adjacent to each other in the frontal lobe of the left or dominant cerebral hemisphere. Conclusions place within the dynamics of a PA cycle that runs through the posterior cortex, the prefrontal cortex, and the environment, and back to the cortex in a circular fashion until the goal is reached. In parallel with that cognitive cycle and interacting with it, a deeper and older cycle processes emotions and instinctual urges through the limbic system. The orbital prefrontal cortex is an integral part of that cycle, which feeds emotional and instinctual influences into the cortex at large. Because reactions to those influences are biologically rooted and weigh heavily on our deci sions, the limbic system constitutes the most primordial evolu tionary root of liberty in the human brain. Liberty, in general, has two major components with oppo site temporal perspective; the first is the ability to choose experi ence from the past, and the second is the ability to choose the future based on the chosen past. The two alternate in tandem with each other under the prefrontal cortex, which integrates past with future at the top of the PA cycle (Chapter 4). The most essentially human is the second, the capacity of the prefrontal cortex to predict events, as well as to select, decide upon, plan, prepare for, and organize goal-directed actions in the immediate or distant future (Chapter 5). Among those actions is spoken and written language, the truly unique patrimony ofour species at the service of all our freedoms and our creative power (Chapter 6). 57